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4. LITHOCARPUS Blume, Bijdr. 526. 1826. 柯属 Ke Shu Pasania Oersted

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4. LITHOCARPUS Blume, Bijdr. 526. 1826. 柯属 Ke Shu Pasania Oersted Flora of China 4: 333–369. 1999. 4. LITHOCARPUS Blume, Bijdr. 526. 1826. 柯属 ke shu Pasania Oersted. Trees or rarely shrubs, evergreen. Winter buds terminal, ovoid to ellipsoid, scales spirally imbricate. Stipules extrapetiolar. Leaves spirally arranged. Inflorescences male, female, or androgynous, in leaf axils toward base of branchlets or in a dense paniculate cluster on subterminal shoots, ± erect. Male inflorescences erect, simple or branched; flowers usually 3–5(–7) in dichasial clusters; perianth 4–6-lobed; stamens 10–12; rudimentary pistil small, enclosed by hairs. Female flowers solitary or in clusters of (2 or)3(–5), 1 or 2(or 3) well developed; perianth 6-lobed; staminodes 10–12; ovary 3(–6) loculed; styles (2 or)3(–5), (0.5–)1–2(–3) mm; stigmas a terminal pore. Cupules grouped together in cymes on rachis but often many aborted, corky, horny, woody, or crustaceous, completely or partly enclosing nut; bracts variously shaped. Nut 1 per cupule. Germination hypogeal; cotyledons flat-convex (although surface between cotyledons may not be completely flat). About 300 species: mainly in Asia, one species in W North America; 123 species (69 endemic) in China. The northern limit of Lithocarpus is on the S flank of the Qinling Mountains. Guangdong, Guangxi, and Yunnan have the highest diversity and the most primitive of the Chinese species. 1a.Nut scar convex (± concave or impressed at margin but conspicuously convex at center in L. cinereus, L. crassifolius, L. handelianus, L. laetus, L. pachyphyllus, and L. variolosus). 2a. Cupules mostly completely enclosing nut. 3a. Scar covering less than 3/4 of nut. 4a. Cupule bracts triangular, inconspicuous, fused with cupule and often apically in 2–4 rings. 5a. Scar covering 1/2–3/4 of nut; leaf blade apex falcate; petiole 1.5–2 cm .................. 14. L. damiaoshanicus 5b. Scar covering ca. 1/4 of nut; leaf blade apex not falcate; petiole less than 1 cm ..................... 15. L. irwinii 4b. Cupule bracts triangular but subulate from middle to apex of cupule, conspicuous, obliquely spreading. 6a. Leaf blade secondary veins 8–12 on each side of midvein; young shoots and petioles with tawny to grayish waxy scalelike trichomes ...................................................................................... 19. L. craibianus 6b. Leaf blade secondary veins more than 12 on each side of midvein; young shoots and leaf blades at least abaxial midvein hairy. 7a. Petiole rarely longer than 1 cm; leaf blade on fruiting branches 9–20 cm wide, secondary veins 21–26 on each side of midvein .................................................................................. 18. L. tephrocarpus 7b. Petiole 2–3 cm; leaf blade on fruiting branches 4–9 cm wide, secondary veins 12–19 on each side of midvein. 8a. Bracts of young cupules subulate, 4-angled; styles with short hairs ......................... 16. L. handelianus 8b. Bracts of young cupules linear, subterete; styles glabrous ............................................. 17. L. amoenus 3b. Scar covering more than 3/4 of nut. 9a. Cupules solitary along rachis .............................................................................................. 13. L. fenzelianus 9b. Cupules in clusters of 3–5 or more, often clusters scattered along rachis. 10a. Cupules entirely covered by bracts. 11a. Cupule bracts spiny; cupules 7–9 cm in diam., including bracts; leaf blade 25–40 cm 1. L. xizangensis 11b. Cupule bracts not spiny; cupules less than 5 cm in diam.; leaf blade rarely to 30 cm. 12a. Cupule bracts straight, multiangular in cross section; wall of nut ca. 4 mm thick ......... 2. L. jenkinsii 12b. Cupule bracts curved, ± round in cross section; wall of nut 2–3 mm thick. 13a. Cupules subglobose; cupule bracts 2–3 mm; nut subglobose to broadly ellipsoid, scar covering 4/5–5/6 of nut .......................................................................................................... 3. L. xylocarpus 13b. Cupules globose; cupule bracts ca. 10 mm; nut turbinate, scar covering ca. 2/3 of nut 4. L. pseudoxizangensis 10b. Cupules sparsely covered by bracts. 14a. Cupule bracts clawlike ........................................................................................................... 5. L. howii 14b. Cupule bracts not clawlike. 15a. Cupules corky or spongy and with smooth concentric lines outside. 16a. Rachis of inflorescences 1–2 cm thick; cupule wall 2–5 mm thick .............................. 6. L. pasania 16b. Rachis of inflorescences 0.6–1 cm thick; cupule wall 5–15 mm thick ....................... 7. L. balansae 15b. Cupules covered with squamose or scurfy bracts and with raised concentric ridges outside. Flora of China 4: 333–369. 1999. 17a. Cupules with bracts conspicuous or inconspicuous, spirally arranged, sometimes scalelike and overlapping. 18a. Cupules flat at apex ......................................................................................... 12. L. lepidocarpus 18b. Cupules rounded at apex ............................................................................... 11. L. amygdalifolius 17b. Cupules with bracts united into 4–8 thin concentric ridges. 19a. Cupules ellipsoid, narrowed to 1 or both ends, 1.2–1.6 cm in diam. ............................ 10. L. levis 19b. Cupules globose to subglobose, 2–3 cm in diam. 20a. Wall of cupule ca. 1.5 mm thick; branchlets of current year blackish when dry .... 8. L. laoticus 20b. Wall of cupule ca. 1 mm thick; branchlets of current year dark brown when dry ..... 9. L. chifui 2b. Cupules mostly not completely enclosing nut. 21a. Wall of cupule more than 3 mm thick or rarely thinner; leaf blade concolorous or with scalelike glands (like tiny drops of water and visible only under high magnifying lens), margin dentate or if entire then abaxially covered with long or stellate hairs. 22a. Nut conspicuously wider than long; cupules ± flat to discoid ............................................ 41. L. pachylepis 22b. Nut ± as long as wide; cupules cupular. 23a. Leaf blade glabrous or with short hairs only along midvein or at axils of secondary veins 40. L. corneus 23b. Leaf blade abaxially hairy. 24a. Hairs simple ................................................................................................................ 38. L. uvariifolius 24b. Hairs stellate or branched. 25a. Leaf blade abaxially densely covered with stellate hairs; secondary veins (15–)20–28 on each side of midvein ........................................................................................................... 39. L. fordianus 25b. Leaf blade sparsely covered with stellate or branched hairs; secondary veins rarely more than 16 on each side of midvein ................................................................................................. 40. L. corneus 21b. Wall of cupule rarely to 2 mm thick; leaf blade not concolorous (concolorous in L. truncatus and sometimes in L. dealbatus) abaxially covered with tightly adherent waxy or lax pulveraceous scalelike trichomes, margin entire or rarely undulate. 26a. Scar usually covering more than 2/3 of nut; cupules enclosing most of nut. 27a. Petiole 3–4 cm ................................................................................................................... 24. L. tabularis 27b. Petiole less than 2.5 cm. 28a. Leaf blade hairy at least abaxially on midvein ........................................................... 20. L. talangensis 28b. Leaf blade glabrous or abaxially covered with tightly adherent waxy scalelike trichomes. 29a. Cupules 3–3.5 cm ........................................................................................................ 21. L. truncatus 29b. Cupules less than 3 cm. 30a. Cupules outside glaucous; bracts imbricate or united into interrupted ring .............. 21. L. truncatus 30b. Cupules outside grayish brown or grayish tawny; bracts (except for subulate tip) fused to cupule, rarely united into concentric rings. 31a. Leaf secondary veins 7–10 on each side of midvein; branches of 2nd- or 3rd-year growth with grayish lamelliform waxy scalelike trichomes ........................................ 22. L. cleistocarpus 31b. Leaf secondary veins 12–16 on each side of midvein; branches without waxy scalelike trichomes ........................................................................................................... 23. L. tenuilimbus 26b. Scar covering at most 1/2 of nut; cupules enclosing 1/3–3/4 of nut. 32a. Nut glabrous. 33a. Petiole 0.2–0.5 cm; leaf blade apex rounded, obtuse, or rarely shortly acute ............. 25. L. crassifolius 33b. Petiole 1–2.5 cm; leaf blade apex acuminate to caudate. 34a. Leaf blade secondary veins conspicuous, abruptly curving, and fusing near margin, secondary and tertiary veins adaxially impressed ................................................................... 26. L. pachyphyllus 34b. Leaf blade secondary veins inconspicuous and not fusing, secondary and tertiary veins adaxially not or only slightly impressed. 35a. Young leaf blades abaxially covered with reddish brown to yellowish brown pulverulous scalelike trichomes ...................................................................................................... 27. L. apricus Flora of China 4: 333–369. 1999. 35b. Young leaf blades abaxially without pulverulous scalelike trichomes. 36a. Leaf buds, young branchlets, and female flower buds without resin; leaf blade abaxially glaucous when dry ............................................................................................... 28. L. variolosus
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